RESEARCH ARTICLE Building a Beetle: How Larval Environment Leads to Adult Performance in a Horned Beetle

The link between the expression of the signals used bymale animals in contests with the traits which determine success in those contests is poorly understood. This is particularly true in holometabolous insects such as horned beetles where signal expression is determined dur-ing metamorphosis and is fixed during adulthood, whereas performance is influenced by post-eclosion feeding. We used path analysis to investigate the relationships between larval and adult nutrition, horn and body size and fitness-related traits such as strength and testes mass in the horned beetle Euoniticellus intermedius. In males weight gain post-eclosion had a central role in determining both testes mass and strength. Weight gain was unaffected by adult nutrition but was strongly correlated with by horn length, itself determined by larval resource availability, indicating strong indirect effects of larval nutrition on the adult beetle’s ability to assimilate food and grow tissues. Female strength was predicted by a simple path diagram where strength was determined by eclosion weight, itself determined by larval nutri-tion: weight gain post-eclosion was not a predictor of strength in this sex. Based on earlier findings we discuss the insulin-like signalling pathway as a possible mechanism by which lar-val nutrition could affect adult weight gain and thence traits such as strength